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Control Stimulation NOTE TO USERS This reproduction is the best copy available. UMI® UNIVERSITY OF CALGARY Raphe Modulation of Circadian Phase by Glenn Yamakawa A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE DEPARTMENT OF PSYCHOLOGY CALGARY, ALBERTA SEPTEMBER, 2009 © Glenn Yamakawa 2009 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 OttawaONK1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-54389-4 Our file Notre reference ISBN: 978-0-494-54389-4 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par Nnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. 1*1 Canada Abstract The midbrain raphe nucleus provides a major input into the SCN, 50% of which contains serotonin. There is mixed evidence as to whether the serotonergic part of this projection is involved in non-photic phase shifting. In order to better characterize the non- serotonergic projections we conducted retrograde tract tracing. Approximately 30% of the projection contained VGLUT3, but not serotonin. To determine if these non- serotonergic projections were important for non-photic phase shifting, the MRN was stimulated in SCN 5-HT lesioned and sham control animals. Intact animals showed a phase advance to midday electrical stimulation of the raphe while the lesioned animals did not. Together, these results show the serotonergic raphe innervation of the SCN is important for non-photic phase shifting and that some of the non-serotonergic raphe to SCN projections may contain glutamate. 11 Acknowledgements I wish to thank my supervisor Dr. Michael Antle, for his patience, insight and allowing me the opportunity to conduct graduate studies in his lab. I wish to thank Dr. Ralph Mistlberger for early opportunities to gain experience in the study of circadian rhythms. I also wish to thank Dr. Cam Teskey, Dr. Dave Glass, Dr. Brian Bland and Dr. John Yeomans for their useful comments and guidance. I wish to thank Glenn Landry for being a mentor and the one who showed me what it took to be in neuroscience research. I also wish to thank my fellow graduate students in the Chronobiology lab, Andrew Brown from the Neural plasticity lab for stimulating discussion and Danica Whalley for technical assistance. Finally, I wish to thank the Natural Science and Engineering Research Council and the Department of Psychology at the University of Calgary for funding this work. m Dedication I wish to dedicate this work to my family for their tremendous love and support. To my mom Cathy for her enthusiasm and strength, and my dad Dick for late night emails and teaching me to find something I am passionate about. IV Table of Contents Abstract ii Acknowledgements iii Dedication iv Table of Contents v List of Figures vii List of Abbreviations viii CHAPTER ONE: GENERAL INTRODUCTION 1 1.1 Master Circadian Pacemaker 1 1.2 Non-Photic Phase Shifting 5 1.2.1 Molecular Effects of Non-Photic Stimuli 8 1.3 The Raphe Complex 10 1.4 Serotonin in the Circadian System 13 1.5 Electrical Stimulation Studies 15 1.6 Serotonin Agonists 15 1.7 Serotonin Antagonists 18 1.8 In Vitro Studies 19 1.9 Evidence Contrary to the Role of 5-HT 22 1.10 Current Studies 23 1.11 Vesicular Glutamate Transporters 24 CHAPTER TWO: RETROGRADE TRACT TRACING 26 2.1 Introduction 26 2.2 Animals and Apparatus 27 2.3 Experiment 1 27 2.3.1 Cholera Toxin p Subunit Ionotophoresis 27 2.3.2 SCN Cholera Toxin Immunocytochemistry 28 2.3.3 5-HT-VGLUT3-Cholera Toxin Immunocytochemistry 29 2.3.4 Analysis 30 2.4 Results 31 2.4.1 Median Raphe and Paramedian Raphe Labelling 33 2.4.2 Dorsal Raphe Labelling 34 2.5 Discussion 35 2.5.1 Retrograde Tract Tracing 35 2.5.2 The Role of VGLUT3 38 2.5.3 Methodological Considerations 42 CHAPTER THREE: MRN STIMULATION 43 3.1 Introduction 44 3.2 Experiment 2 45 3.3 Experiment 2B 46 3.4 Experimental Procedures 47 3.4.1 Perfusion and Immunocytochemistry 48 3.4.2 Analysis 49 3.5 Results 50 v 3.5.1 Behavior during Stimulation 50 3.5.2 Histology 51 3.6 Experiment 2 51 3.6.1 Control Stimulation Procedures 51 3.6.2 Stimulation of the Median Raphe 52 3.7 Experiment 2B 53 3.7.1 Stimulation of the Median Raphe 53 3.8 Discussion 54 3.8.1 What is the Zeitgeber? 54 3.8.2 Circling Behavior 57 3.9 Experimental Design Issues 59 CHAPTER FOUR: GENERAL DISCUSSION 61 4.1 Conclusions 61 4.2 Future Directions 63 4.3 Clinical Importance/Relevance 64 REFERENCES 68 VI List of Figures Figure 2.1 Photomicrograph cases of iontophoresis into the SCN 94 Figure 2.2 Representative triple label histology and mean distribution of cell types retrogradely labelled in the MRN 95 Figure 2.3 Photomicrographs displaying representative examples of retrogradely labelled cell types in the MRN 97 Figure 2.4 Iontophoresis site in the SCN and retrograde labelling in the DRN for one case 98 Figure 2.5 Mean distribution of cell types retrogradely labelling in the DRN 99 Figure 3.1 5-HT labelled histology showing MRN electrode tip placements 101 Figure 3.2 5-HT labelled sections showing intact and lesioned serotonergic input into the SCN 102 Figure 3.3 Mean phase shifts to control and MRN stimulation for experiment 2 .... 103 Figure 3.4 Actograms displaying an animal that received a control manipulation and another that received stimulation of the MRN 105 Figure 3.5 Mean phase shifts to MRN stimulation following cannula injection of vehicle, or 5,7-DHT 106 Figure 3.6 Mean phase shifts to MRN stimulation following vehicle of 5,7-DHT injection for experiment 2B 107 Figure 3.7 Actogram displaying an animal that received a complete lesion of the serotonergic input into the SCN and subsequent stimulation of the MRN 108 vn List of Abbreviations Measurement Terms: Hz hertz Ms millisecond °C degrees Celsius M molar Hrs hours Min minutes Cm centimeters um micrometer uA microampere ml milliliter Other Terms: 5-HT 5-hydroxytryptamine, or serotonin 5-HTR 5-HT transporter 5,7-DHT 5,7-dihydroxytryptamine 8-OH-DPAT 8-hydroxy- 2-(di-n- propylamino) tetralin ABC avidin-biotin complex ANOVA analysis of variance AP anterior/posterior cAMP cyclic adenosine monophosphate CT circadian time CTb cholera toxin beta subunit Cy 2,3,5 Cyanine 2,3, or 5 DAB diaminobenzidine DD constant dark DRN dorsal raphe nucleus DV dorsal/ventral GABA gamma-Aminobutyric acid GHT geniculohypothalamic tract ICC immunocytochemistry IGL intergeniculate leaflet LD light/dark LL constant light ML medial/lateral MRN median raphe nucleus mRNA messenger ribonucleic acid NAAG n-acetyl-aspartyl-glutamate NPY neuropeptide Y PBS phosphate buffered saline Perl period 1 p-ERK phosphorylated extracellular signal responsive kinase PRC phase response curve RHT retinohypothalamic tract Vlll SCN suprachiasmatic nucleus TGF transcription growth factor TH tyrosine hydroxylase VGLUT vesicular glutamate transporter ZT zeitgeber time IX 1 Chapter One: General Introduction 1.1 Master Circadian Pacemaker In today's society of around the clock productivity, trans-meridian travel and shift work it is important to examine ways to function at our best even when our bodies tell us otherwise. Many of the recorded disasters such as the Exxon Valdez, Chernobyl, and Three Mile Island all occurred at times when cognitive and physical functioning was reduced as those involved were on the job during times when they normally would be resting (Folkard and Tucker, 2003). The phenomenon of jet lag is a widespread condition affecting thousands of travelers per year. Jet lag occurs when there is desynchrony between the sleep/wake cycle and the environment that can result in a wide range of symptoms including drowsiness, irritability and gastrointestinal problems (Revell and Eastman, 2005). A similar desynchrony occurs when paramedics, police officers and fire fighters are often called upon to perform complex tasks at night, or in the early morning when they are not operating at their peak efficiency. Recently, it was reported that animals unable to synchronize to the daily light/dark (LD) cycle suffered from extensive cardiovascular and renal damage with dramatically shortened life spans (Martino et al., 2008).
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